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Tylosaurus Kansasensis, a New Species of Tylosaurine (Squamata, Mosasauridae) from the Niobrara Chalk of Western Kansas, USA

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Tylosaurus Kansasensis, a New Species of Tylosaurine (Squamata, Mosasauridae) from the Niobrara Chalk of Western Kansas, USA Netherlands Journal of Geosciences — Geologie en Mijnbouw | 84 - 3 | 231 - 240 | 2005 Tylosaurus kansasensis, a new species of tylosaurine (Squamata, Mosasauridae) from the Niobrara Chalk of western Kansas, USA M.J. Everhart | Sternberg Museum of Natural History, Fort Hays State University, Hays, Kansas 67601, USA. Email: [email protected] Manuscript received: December 2004; accepted: January 2005 Abstract | Tylosaurus kansasensis sp. nov. is described herein on the basis of thirteen specimens collected from the Smoky Hill Chalk (upper Coniacian) of western Kansas, USA. The new species, originally designated Tylosaurus n. sp., co-occurred with T. nepaeohcus and exhibits a number of primitive characters that place it in a basal position in the mosasaur phylogeny. Among the key differences separating this species from other tylosaurines are a shortened, more rounded pre-dental process of the premaxilla, a distinctive quadrate lacking an infrastapedial process, and a parietal foramen located adjacent to the frontal-parietal suture. Keywords: Cretaceous, Niobrara, mosasaur, Tylosaurus kansasensis, Western Interior Sea, new species Introduction was the first mosasaur to be described from Kansas and was originally named 'Macrosaurus' proriger by Cope. The description Mosasaurs (Squamata, Mosasauridae) are extinct marine lizards of the new species by Cope (1869) in the minutes of the June 1 that flourished worldwide during the last 25 million years of meeting of the Academy of Natural Sciences of Philadelphia is the Cretaceous. Mosasaur remains are preserved in marine brief by modern standards, 'He made some remarks on a fine sediments of middle Turanian through late Maastrichtian age, fragment of the muzzle of a large Mosasauroid, which pertained and are especially abundant and well preserved in the Smoky to a cranium of near five feet in length. The pterygoid bones Hill Chalk Member (upper Coniacian - lower Campanian) of the were separated from each other, and support nine teeth. A pecu­ Niobrara Chalk in western Kansas (USA). liarity of physiognomy was produced by the cylindric prolon­ Tylosaurs are among the earliest known mosasaurs from gation of the premaxillary bone beyond the teeth, and a similar the Western Interior Sea of North America. One of the largest flat prolongation of the extremity of the dentary. He referred mosasaur species, Tylosaurus proriger, was first described from the species to Macrosaurus Owen, under the name M. proriger! the upper Smoky Hill Chalk (early Campanian) by Cope (1869). The following year, Cope (1870, pi. 12, figs 22, 23) described The remains of tylosaurine mosasaurs have also been collected the same specimen more completely, figured it and, with little from Europe, Japan, New Zealand and Antarctica. Remains of explanation, referred it to another European genus, Liodon a second, earlier tylosaurine species, Tylosaurus nepaeohcus, Owen. Marsh (1872a, p. 461), apparently recognising significant occur in the lower Smoky Hill Chalk (upper Coniacian) along differences between American and European mosasaurs, with those of Platecarpus tympaniticus and Clidastes liodontus. proposed a new genus {'Rhinosaurus', nose lizard) from a more The type specimen of Tylosaurus proriger (MCZ 4374) was complete specimen (YPM 1268) he had collected 'on the south discovered near Monument Rocks in Gove County, and was side of the Smoky Hill River' in 1871. However, that name was obtained by Professor Louis Agassiz during his 1868 visit to preoccupied and Cope (1872, p. 141) proposed the genus name western Kansas (Cope, 1869; Williston, 1898; Russell, 1967). It Rhamphosaurus. In a brief note, Marsh (1872b, p. 147) wrote Downloaded Netherlandfrom https://www.cambridge.org/cores Journal of Geoscience. IP address:s — Geologi 170.106.202.8e en Mijnbou, on 30 Sepw |2021 84 at- 10:04:103 | 200, subject5 to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0016774600021016 that, 'as this name (Rhinosaurus) proves to be preoccupied, it in a phylogenetic analysis of the Mosasauroidea by Bell (1993, may be replaced with Tylosaurus. The name Rhamphosaurus, 1997) where it was placed as the basal member of the since suggested by Prof. Cope, cannot be retained, as it was Tylosaurine in his informal subfamily 'Russellosaurinae'. given to a genus of lizards in 1843 by Fitzinger.' Leidy (1873, Thirteen specimens of Tylosaurus kansasensis sp. nov. p. 274) was the first to place 'Macrosaurus' proriger Cope, 1869 (Table 2) are here reported from the collections of five museums. into Tylosaurus Marsh. Nine of these specimens, including the holotype and one As an example of the early skirmishes ('Bone Wars') paratype, are curated in the Sternberg Museum of Natural between Cope and Marsh over the naming of newly discovered History, Fort Hays State University, Hays, Kansas. As near as can fossils, Cope (1874, 1875) apparently ignored Marsh's revision, be determined, all of the Kansas specimens documented in this and named a second Tylosaurus species 'Liodon' nepaeolicus report are from the lower one-third (upper Coniacian) of the (AMNH 1565) from remains discovered by B.F. Mudge in the Smoky Hill Chalk Member of the Niobrara Chalk (Appendix 1). lower Smoky Hill Chalk along the Solomon (Nepaholla) River (Everhart, 2002). The genus name went unchallenged until Table 1. Characters of Tylosaurus kansasensis sp. nov. (adapted from Bell, Merriam (1894, p. 24) placed 'Liodon' nepaeolicus Cope, 1874 1993). into Tylosaurus. Regarding Cope's description, Williston (1898, p. 176) wrote that, 'the characters given - such as may be valid 1. Premaxilla rostral foramina large - are altogether too slight to distinguish the species, and I do 2. Infrastapedial process of quadrate greatly reduced or absent not think that T. nepseolicus is entitled to recognition.' Since 3. Medial ridge of quadrate diverges ventrally that time, the second species has been included in work by 4. Keel on dorsal midline of frontal high, thin and well developed Russell (1967), Schumacher (1993), Sheldon (1996), Bell (1993, 5. Medial sutural flanges from frontal extend long distance onto 1997) and Everhart (2002), and is considered here to be valid parietal on the basis of more complete specimens discovered since 6. Parietal foramen adjacent to or invades fronto-parietal suture Williston's time. However, Everhart (2002) noted that besides 7. Dorsal postorbitalfrontal with low rounded transverse ridge the differences in time of occurrence of T. nepaeolicus and 8. Posteroventral angle of jugal virtually 90 degrees T. proriger, only minor morphological features (and the larger 9. Ectopterygoid does not contact maxilla size of T. proriger) appear to separate the two species. Whether 10. Quadrate suprastapedial process without constriction or not T. nepaeolicus and T. proriger represent a single chrono- 11. Quadrate ala thick species is a question that requires further study. 12. Alar concavity of quadrate shallow Russell (1967), Schumacher (1993) and Sheldon (1996) 13. Radius slightly expanded reported that T nepaeolicus is found in the lower chalk (upper Coniacian - lower Santonian), while T. proriger is known only from the upper chalk. In their review of museum collections, Abbreviations neither Schumacher (1993) nor Sheldon (1996) reported specimens of T. proriger from below the upper Santonian. The AMNH - American Museum of Natural History, New York, NY; earliest known remains of T. proriger (FFHM 1997-10) were FHSM - Fort Hays State University, Sternberg Museum of described by Everhart & Everhart (1997) from the middle Natural History, Hays, Kansas; FGM - Fryxell Geology Museum, Santonian chalk of Gove County. In his revision of mosasaur Augustana College, Rock Island, Illinois; LACMNH - Los biostratigraphy of the Smoky Hill Chalk, Everhart (2001) noted Angeles County Museum of Natural History, Los Angeles, CA; that T. nepaeolicus was present in the upper Coniacian and MCZ - Museum of Comparative Zoology, Harvard University, lower Santonian chalk in Kansas. Cambridge, MA; TMM - Texas Memorial Museum, University of While it is possible that Cope (1875, pi. 28, figs 8, 9) may Texas at Austin, TX; YPM - Peabody Museum of Natural History, have figured the distinctive, abbreviated premaxilla of the Yale University, New Haven, CT. third species as that of 'Liodon dyspelor', Stewart (1990) was the first to document the existence of Tylosaurus n. sp. An I Systematic palaeontology examination of the extensive 1870s collection from Kansas in the Yale Peabody Museum by Bell (1993) determined that a Order Squamata specimen of the third species (YPM 40796) had been collected Family Mosasauridae in 1876, but was then identified no further than Tylosaurus sp. Genus Tylosaurus Marsh, 1872b The characteristics of an 'undescribed' species of Tylosaurus were discussed by Schumacher (1993) in regard to specimens in I Tylosaurus kansasensis sp. nov., Figs 1 - 9 the collection of the Sternberg Museum (FHSM VP-78, VP-2295 and VP-3366), and a list of diagnostic characters (Table 1) was Material - Holotype is FHSM VP-2295, a complete, articulated compiled by Bell (1993). The undescribed species was included skull and seven cervical vertebrae, preserved in dorsal/left Downloaded from https://www.cambridge.org/core. IP address: 170.106.202.8, on 30 Sep 2021Netherland at 10:04:10s ,Journa subject lto o thef Geosdence Cambridge Cores — terms Geologi of use,e e availablen Mijnbou at w | 84 - 3 | 2005 https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0016774600021016
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